the amphibians and reptiles of the lore lindu national...

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Herpetofauna of Lore Lindu National Park area

All articles available online at http://www.salamandra-journal.com© 2011 Deutsche Gesellschaft für Herpetologie und Terrarienkunde e.V. (DGHT), Rheinbach, Germany

SALAmANDRA 47(1) 17–29 20 February 2011 ISSN 0036–3375

The amphibians and reptiles of the Lore Lindu National Park area, Central Sulawesi, Indonesia

Thomas Cherico Wanger1,2, Iris motzke2, Shahabuddin Saleh3 & Djoko T. Iskandar4

1) Environment Institute and School of Earth and Environmental Sciences, University of Adelaide, SA 5095, Australia2) Agroecology, University of Göttingen, 37073 Göttingen, Germany

3) Agrotechnology, Faculty of Agriculture, University of Tadulako Palu, Indonesia4) School of Life Sciences and Technology, Institut Teknologi Bandung, 40132-Bandung, Indonesia

Corresponding author: Thomas Cherico Wanger, e-mail: [email protected]

manuscript received: 03 April 2010

Abstract. While land-use change is rapid throughout Southeast Asia, the island of Sulawesi (Indonesia) is of pressing con-servation concern because of its exceptional number of endemic species. However, a lack of good identification literature for certain taxa such as amphibians and reptiles (apart from snakes) substantially delays ecological research in this region. Here, we compile an illustrated species list based on three years of research in and around the Lore Lindu National Park (LLNP) area and supplement it with data from the literature. In total, our survey and the literature review revealed 25 am-phibian and 54 reptile species in five and thirteen families, respectively. Our results highlight the LLNP area as an impor-tant herpetological endemism hotspot in the region. Appropriate utilization of species lists like this may facilitate capacity-building of local scientists and provide a knowledge base for local guides working in ecotourism.

Key words. Biodiversity, capacity-building, conservation, ecotourism, species list, Southeast Asia, Sulawesi.

Abstrak. Seiring dengan cepatnya konversi lahan di Asia Tenggara, Sulawesi (Indonesia) menjadi kawasan yang sangat penting untuk dikonservasi atas dasar tingginya keaneka-ragaman jenis, terutama untuk jenis yang endemik. Walaupun demikian, ketidak tersediaan kepustakaan yang mutakhir untuk pengenalan jenis (kecuali untuk jenis-jenis ular) menjadi penghambat utama dalam penelitian ekologi di daerah ini. Untuk tujuan tersebut, makalah ini dilengkapi dengan gambar yang dihasilkan dari penelitian selama tiga tahun di dalam kawasan Taman Nasional Lore Lindu maupun daerah sekitarnya dan ditambah dengan informasi dari kepustakaan yang ada. Berdasarkan informasi yang telah dihimpun, 25 jenis amfibi dan 54 jenis reptil dijumpai di dalam kawasan Taman Nasional Lore Lindu, mewakili lima famili katak dan 13 famili rep-til. Hasil ini menunjukkan bahwa TNLL merupakan kawasan endemik yang penting untuk Sulawesi. Diharapkan maka-lah ini dapat bermanfaat untuk memberikan data dasar bagi masyarakat setempat dan juga berguna untuk kepentingan ekoturisme.

Introduction

Based on deforestation rates, climate change predictions, and the extent of current land-use change, Southeast Asia is a region of high conservation priority (Sodhi et al. 2010a, Brooks et al. 2006). However, ecological research efforts and suitable protection of species naturally depend on the knowledge of species occurrences in a particular area. This information in readily accessible form is lack-ing for several taxonomic groups and regions in Southeast Asia (Sodhi and Brook 2006). In order to facilitate argu-ments to policy makers for the protection and preservation of invaluable conservation areas and informed ecotourism (i.e., knowledgeable local guides), such accessible data is of crucial importance.

The island of Sulawesi (Indonesia) is a hotspot of en-demism even on a global scale (Whitten et al. 2002), but knowledge about herpetological diversity and identifi-cation literature are both very limited (but see de Lang and Vogel 2005 for snakes). This is a worrying tendency

as these two taxonomic groups contain many of the most threatened vertebrates on the planet (IUCN 2009, http://www.iucnredlist.org) and the ecological impacts of land-use change urgently need to be better understood (Sodhi et al. 2010b). Based on three years of collecting survey data and the available literature, we here provide the first her-petological species list with IUCN threat status and infor-mation on abundance, habitat use, and maximum eleva-tion range, supplemented with pictures for most species encountered in the Lore Lindu National Park, LLNP, area in Central Sulawesi province. We discuss how studies like this may greatly benefit capacity-building of local scientists and tourism in the LLNP region, one of the largest rainfor-est national parks in Indonesia.

Material and methods

We collected our data in and around the LLNP, situated south of Palu, the capital of Central Sulawesi province (Fig.

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Thomas Cherico Wanger et al.

1). The park covers 230,000 ha of mostly pristine tropical rainforest that partly covers mountain ranges with a maxi-mum altitude of 2356 m above sea level (mount Nokila-laki). The surroundings are dominated by agricultural land-use, mainly for cacao and coconut plantations, as well as rice cultivation. Native forest diversity is highly threat-ened by hunting activities and the land-use change of pris-tine forest into agricultural plantations.

We sampled amphibians and reptiles on 110 m transects with visual and acoustic encounter surveys (Heyer et al. 1994) and a 25-minute time constraint. more than 100 transects in six habitat types (primary and secondary rain-forest, two cacao agroforest types, rice paddies, and open areas) were checked up to six times (three times during the day and night each). Our sampling from December 2007 until October 2009 covered wet and dry seasons and an el-evational range from 400 to 2300 m above sea level (mount Nokilalaki).

In addition to our sampling, the species list including abundance, habitat use, and elevation ranges was com-

plemented with data from the literature (for references see Tabs. 1, 2). IUCN status was obtained from the IUCN RedList database (IUCN 2009, http://www.iucnredlist.org).

Nomenclature used here follows Frost (2009) for am-phibians and mcDiarmid et al. (1999) and de Lang and Vogel (2005) for snakes. For other reptiles, we used the literature sources cited in Table 2.

Results

We identified 25 amphibian species in five families and 54 reptile species in 13 families (Tab. 1). Reptiles comprised 22 snake, 28 lizard, and four turtle species (Tab. 2). While 32.0 % of all amphibian species found are not yet classified in the IUCN Red List, only four of the reptile species (the turtles) have been assessed until now. Our elevation data suggests that four of the new Limnonectes species are po-tentially high-elevation specialists and one rhacophorid is

Figure 1. map of the Lore Lindu National Park (LLNP) area in Sulawesi at three different scales: within Southeast Asia (A); in Sulawesi (B); in Central Sulawesi (C). The map shows the mountainous terrain of Central Sulawesi with the two highest peaks, both inside (mount Nokilalaki, 2357 m, indicated by a white dot) and outside the LLNP (mount Rore Katimbu 2610 m, indicted by a star). Land-use change from pristine forest into agricultural areas is evident all around the park, but most prominently in the northern and eastern sections. The park boundary is indicated by a white line; the arrow in (C) indicates north for (A) to (C).

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Amphibian species IUCN Red List

Specialist Habitat encountered

Abun-dance

Source LLNP Elevation Source El

BufonidaeIngerophrynus biporcatus LC N O Iskandar & Tjan

1996<1000 Sodhi et al. 2008

Duttaphrynus melanosticus LC N AF, CP, HS C Wanger et al. 2009, This study

<1800 Sodhi et al. 2008

Ingerophrynus celebensis LC N PF, SF, AF, CP, OA

O Wanger et al. 2010a, Wanger et al. 2009

<1000 Sodhi et al. 2008

MicrohylidaeCallulops sp. NE Y PF R This study F1200 This studyKaloula baleata LC N C Amphibia Web 2009 <800 Sodhi et al. 2008;

updated from our studyKaloula pulchra LC N PF C Wanger et al. 2010a F800 This studyOreophryne sp. 1 NE N PF, SF R Wanger et al. 2010a F800 This studyOreophryne sp. 2 NE Y PF R This study F800 This studyRanidaeHylarana celebensis LC N PF, SF C Wanger et al. 2010a,

Wanger et al. 2009<800 Sodhi et al. 2008;

updated from our studyHylarana erythraea LC N C Amphibia Web 2009 <1200 Sodhi et al. 2008 Hylarana macrops NT Y PF, SF R Amphibia Web 2009,

This study<1000 Sodhi et al. 2008

Hylarana mocquardii LC N PF, SF, AF, CP, OA

C Inger et al. 2009, This study

<1000 Sodhi et al. 2008

DicroglossidaeFejervarya cancrivora LC N PF, SF, AF, CP,

OA, RFC This study <400 Sodhi et al. 2008;

updated from our studyFejervarya limnocharis LC N PF, SF, AF, CP,

OA, RFC This study <2000 Sodhi et al. 2008

Limnonectes modestus LC N C Amphibia Web 2009 <1550 Sodhi et al. 2008; updated from our study

Limnonectes cf. arathooni NE Y PF R Wanger et al. 2010a, Wanger et al. 2009

F2300 This study

Limnonectes cf. modestus NE N PF, SF R Wanger et al. 2010a, Wanger et al. 2009

F1400 This study

Limnonectes cf. heinrichi NE Y AF R Wanger et al. 2010a F1600 This studyLimnonectes sp. (medium 3) NE Y PF R Wanger et al. 2009 F1550 This studyOccidozyga celebensis LC N C Amphibia Web 2009 <1550 Sodhi et al. 2008;

updated from our studyOccidozyga semipalmata LC Y PF C This study <1550 Sodhi et al. 2008;

updated from our studyRhacophoridaePolypedates leucomystax LC N PF, SF, AF, CP,

OAC Amphibia Web 2009,

This study<1550 Sodhi et al. 2008

Rhacophorus edentulus DD ? ? Iskandar & Tjan 1996

<1000 Iskandar & Tjan 1996

Rhacophorus monticola NT Y C Amphibia Web 2009 >1000 Sodhi et al. 2008 Rhacophorus sp. NE ? PF, SF C J.A. mcGuire, pers.

comm.<300 Iskandar & Tjan

1996; this study

Table 1. Amphibian species of the Lore Lindu National Park. Red List classifications follow the criteria of IUCN (IUCN 2009, http://www.iucnredlist.org): CE = Critically Endangered; E = Endangered; V = Vulnerable; NT = Near Threatened; LC = Least Concern; DD = Data Deficient; NE = not evaluated by the IUCN; Specialist [Y/N/?] = species considered a pristine-forest specialist/disturbance-tolerant species/not known; Habitat encountered = habitat where we encountered the species during the three-year survey; abbreviations used: PF = Rainforest; SF = secondary forest; AF = Cacao agroforest; CP = Cacao plantation; OA = Open area; RF = Rice paddy. Abundance was classified as C = commonly encountered; R = rare (as mentioned in the literature). Source LLNP = literature source that lists the species for Central Sulawesi Province/Lore Lindu National Park. Note that the studies by Wanger et al. (2009, 2010a) employed the same sampling methods and were conducted over the same time period as this study. Data from Amphibia Web (2009, http://amphibi-aweb.org/). Elevation = maximum elevation range in metres above sea level given in the Source El reference. If no elevation data could be found in the literature, we present the elevation at which we found the species (indicated with “F” before the figure). The symbols < and > denote occurrence below or above the elevation given, respectively. The word “lowland” is used when it was mentioned that the species would occur in the lowland without giving an elevation in metres. NA = information on the elevation range is not available.

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Table 2. Reptile species of the Lore Lindu National Park. For a detailed column header and explanation of abbreviations used see legend of Tab.1. Further abbreviations: + D.T. Iskandar noted that Rhabdophis chrysargoides does not occur in Sulawesi, despite the record in de Lang & Vogel (2005); ++ we noted indications that shops were selling specimens collected in the Lore Lindu area; this was confirmed by locals. Note that the works by Wanger et al. (2009, 2010a) were also based on this three-year sampling survey and consequently employed the same sampling methods. $: contains a key to the Cyrtodactylus species of Sulawesi.

IUCN Red List

Specia-list

Habitat encountered

Abun-dance


Snake speciesColubridaeAhaetulla prasina NE N SF, AF, CP C Wanger et al. 2009 < 1300 deLang & Vogel 2005Boiga dendrophila gemmicincta NE Y C deLang & Vogel 2005 Lowland de Rooij 1917Boiga irregularis NE N AF C Wanger et al. 2010a < 1400 deLang & Vogel 2005Calamaria nuchalis NE N C deLang & Vogel 2005 < 610 de Rooij 1917Calamaria sp. NE N SF C This study F800 This studyCalamaria virgulata NE N C deLang & Vogel 2005 NA deLang & Vogel 2005Chrysopelea paradisi celebensis NE N C deLang & Vogel 2005 < 1300 deLang & Vogel 2005Dendrelaphis pictus pictus NE N AF, CP C Wanger et al. 2009 < 1400 deLang & Vogel 2005Elaphe erythrura celebensis NE N C deLang & Vogel 2005 < 1100 deLang & Vogel 2005Gonyosoma jansenii NE N R deLang & Vogel 2005 < 1000 deLang & Vogel 2005Lycodon stormi NE Y PF, SF, AF R This study F820 This studyOligodon waandersi NE Y C deLang & Vogel 2005 < 1200 deLang & Vogel 2005Rabdion forsteni NE N SF C This study < 1830 deLang & Vogel 2005Rhabdophis callistus NE N SF R Wanger et al. 2009,

Wanger et al. 2010a, +< 1200 This study

Psammodynastes pulverulentus pulverulentus

NE N SF C Wanger et al. 2010a < 2000 deLang & Vogel 2005

Ptyas dipsas NE N PF, SF, AF, CP, OA

R This study < 1100 deLang & Vogel 2005

Xenochrophis trianguligerus NE N PF, SF, AF, CP, OA, RF

C This study < 1400 deLang & Vogel 2005

CylindrophiidaeCylindrophis melanotus NE Y C deLang & Vogel 2005 < 1200 deLang & Vogel 2005ElapidaeOphiophagus hannah NE N PF, SF, AF,

CP, OA, RFR This study; ++ < 1800 deLang & Vogel 2005

PythonidaePython reticulatus reticulatus NE N C deLang & Vogel 2005 < 1300 deLang & Vogel 2005ViperidaeTropidolaemus subannulatus NE N C deLang & Vogel 2005;

Vogel et al. 2007< 1300 deLang & Vogel 2005

XenopeltidaeXenopeltis unicolor NE N AF, CP C Wanger et al. 2009,

Wanger et al. 2010a< 1300 deLang & Vogel 2005

Lizard speciesAgamidaeBronchocela celebensis NE N PF, SF,AF, CP C Wanger et al. 2009 < 1200 This studyBronchocela cristatella NE N PF, SF, SF, CP C This study < 1600 manthey &

Grossmann 1997Draco spilonotus NE ? ? mcGuire et al. 2007 < 1016 mcGuire et al. 2007Draco walkeri NE ? ? mcGuire et al. 2007 < 1840 mcGuire et al. 2007GekkonidaeCyrtodactylus spinosus NE ? ? ? Wanger et al. 2009 ~ 600 Linkem et al. 2008Cyrtodactylus wallacei NE ? ? Hayden et al. 2008, $ NA Hayden et al. 2008Cyrtodactylus jellesmae NE Y SF, AF R Wanger et al. 2009 < 850 Wanger et al. 2009Gekko gecko NE Y CP, OA C This study < 900 manthey &

Grossmann 1997Gekko monarchus NE N PF, SF,AF, CP,

OAO This study < 1500 manthey &

Grossmann 1997

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adapted to lower elevations. The remaining amphibians oc-cur in the lowland or within a broad elevation range. None of the reptiles seem to be high-elevation specialists. For pictures of most of the encountered amphibian and lizard species see the Appendices. Photographs of most snakes of Sulawesi can be found in de Lang and Vogel (2005).

Discussion

Due to its geological past, Sulawesi’s herpetofaunal diver-sity is lower compared to the neighbouring islands such as Borneo, the moluccas or Papua (e.g., Whitten et al. 2002).

Being a global endemism hotspot, our species list supports the notion of Iskandar and Tjan (1996) that Central Su-lawesi is a very valuable area for the herpetofauna of the island. However, many areas in the region have not yet been covered by herpetological surveys – and the fact that dozens of species are awaiting formal description suggests that many more are likely to be discovered (D.T. Iskandar, pers. obs.). Our current result of few high-elevation spe-cialist amphibian and no such reptile species may just re-flect an under-sampling of mountainous habitats. A better estimate of an approximate number of elevation specialists is crucial, however, because these species will be most se-verely affected by climate change.

IUCN Red List

Specia-list

Habitat encountered

Abun-dance


Gehyra mutilata NE N C Whitten et al. 2002 < 1500 manthey & Grossmann 1997

Hemidactylus frenatus NE N OA C This study < 1600 manthey & Grossmann 1997

Hemidactylus platyurus NE N C Iskandar & Tjan 1996 < 1200 de Rooij 1915ScincidaeSphenomorphus celebense ? ? ? Iskandar & Tjan 1996 < 1200 de Rooij 1915Emoia atrocostata NE Y O Iskandar & Tjan 1996 Lowland de Rooij 1915Emoia caeruleocauda NE Y C Iskandar & Tjan 1996 < 920 de Rooij 1915Eutropis sp. NE N SF, AF, CP,

OAC Wanger et al. 2009,

Wanger et al. 2010a< 900 This study

Eutropis multifasciatus NE Y CP, OA C Wanger et al. 2009, Wanger et al. 2010a

< 1800 manthey & Grossmann 1997

Eutropis rudis NE N PF, SF, AF, CP

C Wanger et al. 2009, Wanger et al. 2010a

< 900 This study

Lamprolepis smaragdina NE ? ? Iskandar & Tjan 1996 < 300 de Rooij 1915Parvoscincus sp. NE ? PF, SF, AF,

CP? Wanger et al. 2009,


Lipinia inconspicua NE ? ? ? Wanger et al. 2009, Wanger et al. 2010a

< 1200 de Rooij 1915

Sphenomorphus cf. textus NE ? PF, AF, CP, OA

? Wanger et al. 2009, this study

< 900 This study

Sphenomorphus nigrilabris NE Y PF, SF, AF, CP

? Wanger et al. 2009, Wanger et al. 2010a

< 900 This study

Sphenomorphus tropidonotus NE ? ? Iskandar & Tjan 1996 < 900 This studySphenomorphus cf. variegatus NE N PF, SF, AF,

OA? Wanger et al. 2009,


Tropidophorus baconi NE ? ? Hikida et al. 2003 < 1000 This studyDibamidaeDibamus celebensis NE ? R Iskandar & Tjan 1996 < 1400 de Rooij 1915VaranidaeVaranus salvator NE N C Iskandar & Tjan 1996,

this study< 1200 mcKay 2006

Turtle speciesGeoemydidaeCuora amboinensis V N C D.T. Iskandar, pers.

observ.Lowland manthey &

Grossmann 1997Leucocephalon yuwonoi CE ? R Ives et al. 2008 Lowland D.T. Iskandar, pers.

observ.TestudinidaeIndotestudo forstenii E ? R Ives et al. 2008 Lowland de Rooij 1915TrionychidaeAmyda cartilagínea V ? ? ? Koch et al. 2008 Lowland de Rooij 1915

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During this survey, we did not encounter some of the species found during previous surveys. One reason for this could be that species have declined in numbers or became locally extinct due to the rapidly progressing deforestation. This development is evident in the northeastern parts of the park (the Dongi-Dongi area), where immigrants from South Sulawesi, but also other parts of Indonesia, rapid-ly convert the forest into cacao plantations (Weber et al. 2007). As, for example, one of the study areas of the cited studies (Iskandar & Tjan 1996) focused in parts on this particular site, several species are likely to have since be-come at least locally extinct. In addition, the present study has not exhaustively sampled the area, even though most relevant habitat types were covered. We did not use pitfall traps because the effort required to maintain these setups are cost- and labour-intensive, but may not trade off well in terms of what they add to the species list. The penalty is that we are likely to have missed several secretive and fos-sorial species such as Dibamus spp., Calamaria spp., Pseu-dorabdion spp., and small skinks. The high herpetological diversity we found within this endemic hotspot is, how-ever, severely threatened by the rapid forest conversion for cacao plantation; already one million ha of Sulawesi’s land surface have been converted into these plantations to serve the global demand for cocoa (Direktorat Jenderal Perke-bunan 2008).

Recently, two ecological studies explicitly investigated the effects of land-use change on amphibians and reptiles in the Lore Lindu National Park(Wanger et al. 2009, 2010). Results indicated clearly that amphibians will be severely impacted by forest conversion, most likely due to their sen-sitivity to desiccation. For these animals, extensive pristine forest coverage is crucial for survival. many reptiles, how-ever, may benefit from agricultural habitats, as their life histories in general are more dependent on thermoregu-lation in open canopy spots (Huey et al. 2009, Wanger et al. 2010a). The sensitivity of both groups to climatic al-terations (Pounds et al. 2006, Huey et al. 2009, Wanger et al. 2009) can – to a certain extent – be made allowance for by a considerate structural management of these agri-cultural habitats. Cacao plantations with sufficient canopy cover, leaf litter thickness, and heaps of branches have a likely potential for sustaining greater herpetofaunal spe-cies richness and abundance than heavy-handedly man-aged, cleared lands. However, the main beneficiaries will certainly be common and “robust” species such as the Celebes toad (Ingerophrynus celebensis), the common sun skink (Eutropis multifasciatus), and a common but as yet undescribed skink species (Eutropis sp.).

Besides supporting conservation arguments with data (i.e., giving rough figures of how many species occur in an area), such species lists may greatly facilitate capacity-building of local researchers and field guides for ecotour-ism. For local researchers, learning the members of a local fauna by their scientific names is an important means for conducting sound scientific projects. moreover, if ecotour-ists come to remote places to see wildlife, amphibians are often the only vertebrates they get to see. It is beneficial if local guides can then name a ‘Celebes Toad’ on the basis of a brochure instead of referring to it as ‘Tete’ (i.e., the spe-cies’ name in the local language [Bahasa Toro]). Booklets for tourists will in addition provide financial support for

the national park bureau to fund the paying of often insuf-ficient salaries for local rangers.

Future research efforts should target an extensive and complete coverage of the herpetofauna of Sulawesi (D.T. Iskandar et auct., unpubl.) based on genetic and mor-phological analyses of species, many of which have so far remained undescribed. Simultaneously, explorative sur-veys in the region should continue in particular in re-mote mountainous areas where no herpetofaunistic sam-pling has so far been conducted. more ecological studies are needed to assess the effects of pesticides on amphibians and reptiles as well as biodiversity in general as forest is converted into plantation habitats (Wanger et al. 2010b). Conservation value assessments may provide guidelines of how to make secondary habitats more hospitable to biodi-versity. However, this will only work if we can take poten-tially detrimental pesticide effects into account.

Acknowledgements

We would like to thank our assistants in Sulawesi, who made this project possible and in particular, the people of Toro for their hospitality and support. We thank S. Howard and U. manthey for providing comments on the species list and G. Gillespie, S. Howard, J. A. mcGuire and U. manthey for giving us per-mission to use their photographs. Surveys were conducted un-der the research permits from Kementerian Negara Riset dan Te-knologi Republik Indonesia (RISTEK; #1899/FRP/Sm/VIII/2008; to T.C. Wanger) and Lembaga Ilmu Pengetahuan Indonesia (LIPI; #7374a/SU/KS/2007; to T.C. Wanger). Funding was pro-vided through the German Science Foundation (DFG), an En-deavour International Postgraduate Research Scholarship, and a University of Adelaide Scholarship (to T.C. Wanger).

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Appendix 1. Agamidae: A = Bronchocela celebensis; Gekkonidae: B = Gekko gecko (West malaysia; Photo by U. manthey); C = Cyrto-dactylus jellesmae; D = Cyrtodactylus spinosus (reproduced with permission from the authors and Allen Press); E = Cyrtodactylus wal-lacei (reproduced with permission from the authors and Allen Press); F = Gekko monarchus (West malaysia; Photo by U. manthey); G = Gehyra mutilata (Photo by G. Gillespie); H = Hemidactylus frenatus (Thailand; Photo by U. manthey).

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Appendix 2. Gekkonidae: A = Hemidactylus platyurus (Thailand; Photo by U. manthey); Scincidae: B = Emoia atrocostata (Photo by G. Gillespie); C = Eutropis n. sp.; D = Eutropis multifasciatus; E = Eutropis rudis; F = Parvoscincus sp.; G = Sphenomorphus cf. textus; H = Sphenomorphus nigrilabris.

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Appendix 3. Scincidae: A = Sphenomorphus tropidonotus (Photo by G. Gillespie); B = Sphenomorphus cf. variegatus; C = Tropidopho-rus baconi (Photo by J. A. mcGuire); D = Lamprolepis smaragdina (Photo by S. Howard); Dibamidae: E = Dibamus celebensis (Photo by G. Gillespie); Varanidae: F = Varanus salvator; Agamidae (both pictures reproduced with permission from the authors and Allen Press): G = Draco walkeri; H = Draco spilonotus.

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Appendix 4. Bufonidae: A = Duttaphrynus melanostictus; B = Ingerophrynus celebensis (juvenile); microhylidae: C = Callulops n. sp.; D = Kaloula baleata; E = Kaloula pulchra (West malaysia; Photo by U. manthey); F = Oreophryne n. sp.1; G = Oreophryne n. sp.2; Di-croglossidae: Fejervarya cancrivora

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Appendix 5. Dicroglossidae: A = Limnonectes cf. modestus; B = Limnonectes cf. arathooni; C = Limnonectes cf. heinrichi; D = Fejervarya limnocharis; E = Occidozyga semipalmata; Ranidae: F = Hylarana celebensis; G = Hylarana erythraea; H = Hy-larana macrops.

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Appendix 6. Ranidae: A = Hylarana mocquardii; Rhacophoridae: B = Polypedates leucomystax; C = Rhacophorus sp. (Photo by J. A. mcGuire)

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